Unified phylogenetic species concept: taking subspecies and race out of science: postmodern theory applied to the Potamophylax cingulatus group (Trichoptera, Limnephilidae)

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Opusc. Zool. Budapest, 2018, 49(1): 33–70

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Unified phylogenetic species concept: taking subspecies and race out of science: postmodern theory applied to the Potamophylax

cingulatus group (Trichoptera, Limnephilidae)

J.OLÁH¹,T.ANDERSEN²,S.BESHKOV³,C.CIUBUC⁴,G.COPPA⁵,H.IBRAHIMI⁶,T.KOVÁCS⁷, J.OLÁH⁸ JR.&B.SZCZESNY⁹

1János Oláh, Residence postal address: Tarján u. 28, H-4032 Debrecen, Hungary. E-mail: profolah@gmail.com

2Trond Andersen, Department of Natural History, University Museum of Bergen, University of Bergen, P.O. Box 7800, N-5020 Bergen, Norway. E-mail: trond.andersen@um.uib.no

3Stoyan Beshkov, National Museum of Natural History, 1 Tsar Osvoboditel Blvd. 1000 Sofia, Bulgaria.

E-mail: beshkov@nmnhs.com

4Constantin Ciubuc, Sinaia Zoological Research Station, University of Bucharest, Cumpatu 5, Sinaia, R-106100, Romania. E-mail: ciubuc1206@gmail.com

5Gennaro Coppa, 1, rue du Courlis, F-08350 Villers-sur-Bar, France. E-mail: gennaro.coppa@wanadoo.fr

6Halil Ibrahimi, University of Prishtina, Faculty of Mathematics and Natural Sciences, Department of Biology, Mother Teresa p.n., 10000 Prishtina, Kosovo. E-mail: halilibrahimi@yahoo.com

7Tibor Kovács, HNHM Mátra Museum, Kossuth Lajos u. 40, H-3200 Gyöngyös, Hungar. kaoti@t-online.hu

8János Oláh jr., Residence postal address: Tarján u. 6, H-4032 Debrecen, Hungary.

E-mail: sakertour@gmail.com

9Bronislaw Szczesny, Institute of Nature Conservation, Polish Academy of Sciences, Kraków, Poland.

E-mail: szczesny@iop.krakow.pl

Abstract. The subspecies of the biological species concept with incomplete reproductive isolation versus the incipient sibling species of the phylogenetic species concept with permeable reproductive barrier are still applied side by side in the everyday practice of taxonomy. Both terms refer to the same organisms diverged mostly in allopatry with various stages of reproductive isolation. Question remained: how human ranks these entities organised by nature? The reliable ranking of living hierarchies is retarded and even obscured by the suppressed state of taxonomy. Disappointing scenario: the science of biodiversity is stuck in century old macromorphologies without innovation of fine phenomics and without exploring its high- tech and high-throughput potential. The empirical science of taxonomy is “modernised” by the neutral DNA marker industry diverting the epistemological focus from empirical to virtual. Virtuality of noumenon is used to camouflage the phenomenon of the adverse environmental processes, the wasteful byproducts of the profit oriented liberalized economy. The sensual reality of species and the accelerated species extinction is effectively masked by the virtual sciences of the abstract: numbers, data, statistics, algorithms, equations, models and ideas. To understand the birth of a young incipient species we have briefly reviewed the postmodern development of the unified phylogenetic species concept. (1) The reality of species and higher phylogenetic taxa. (2) The biological and phylogenetic species. (3) How to delineate phylogenetic species? (4) The infinite versus finite division of phylogenetic species. (5) The construct of the unified species concept. (6) Taking subspecies and race out of science. Without recognition of incipient siblings of the phylogenetic species the biodiversity remains underestimated and the pharisaic anti-science ranking of humans remains with us. The discovery of speciation trait that is the sexual adaptive structures in reproductive barrier building, which are detectable by fine phenomics, gives perspective to find the finite division, the dynamic initial split in the continuous process of diversification. The speciation traits produced by integrative organisation, as opposed to competitive selection, help to unify the operational criteria of the biological species concept that is the speciation by reproductive isolation with the general concept of phylogenetic species that is the causal process of the separately evolving metapopulation lineages. The subspecies and racial ranking is untenable anymore, we suggest taking subspecies and race out of science: the finite division of the initial split detected by speciation traits is the birth of the phylogenetic incipient sibling species. There is no “subspecies”and “races”, as there is no “subindividual” in the biological organisation. In the present caddisfly taxonomy the subspecies remained as a valid status in the Potamophylax

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Oláh et al.: Unified phylogenetic species concept applied to the Potamophylax cingulatus species group

cingulatus caddisfly species group. With a clear distinction between the neutral and adaptive traits in the P. cingulatus species group and applying the subtle and stable shape divergences in phallic fine structures we have proposed to change the taxonomic status of subspecies to incipient sibling phylogenetic species rank: Potamophylax alpinus stat. nov., P. depilis stat nov., P. ibericus stat. nov., P. inermis stat. nov., reinstated the species status of P. cingulatus stat. restit. and we have described three new species: P. fesus Oláh, P. portugalicus Oláh et Szczesny, and P. transalpinus Oláh & Coppa, spp. nov.

Keywords. Suppressed taxonomy, speciation trait, race, subspecies, phylogenetic species, sibling species, Potamophylax cingulatus species group, new species.

INTRODUCTION

axonomy is staggering today in postdarwi- nian contradictions remained non-harmonised as regards the ever-lasting conflict between epis- temology of ranking and ontology of organisational hierarchies: that is (1) how human ranks entities and (2) how entities are organised in nature. What are species, subspecies or races as organised by nature and as ranked by science? In spite of these indispensable questions waiting to be understood, our taxonomy that is the basic empirical and integrative discipline of natural history for answering such questions appears more and more as a neglected and almost dying science.

Why taxonomy is suppressed? Is there any cultural interest or social context not to answer these questions, especially the last one: what are races?

This happens in the middle of the biodiversity cri- sis amplified by global warming. Today all the achievements of high-tech and high-throughput potential of the fine phenomics, the empirical fu- ture of taxonomy, is repressed and retarded by the over financed blind neutral DNA marker industry.

Taxonomists realise their backyard position every day in the western culture: there is no sound genuine taxonomic project possible to launch without at least one component of the modern slogans fabricated in masking industries: DNA sequenc- ing, warming models or evolutionary theories.

Is taxonomy suppressed?

The painful result of this desperate state is clearly documented by the simple fact that “modern” taxonomy, at least our caddisfly taxonomy, is based and practiced primarily on the century old procedures of macromorphologies. The species descriptions and drawings of the Russian sci- entist Martynov (1909, 1915) are still comparable

to, or even exceed the quality of many of our present-day drawings and structural understand- ings. This stagnant condition in taxonomy was created and maintained by non-taxonomists and by distracting movements. In the last eighty years the “modernization” of taxonomy was focused by highly speculative models of mathematicians (Haldane, Fischer, and Wright) and by virtual molecular approaches manifested in neutral DNA sequences of geneticists (Dobzhansky). Virtual artefacts of speciation processes, taxon ranking and species delineations are further deformed by dictates of ideological and political projects generated in the dominating practices of Darwinism.

This kind of “modernization” is getting more transparent today as an intentional movement to replace and to divert the focus from empirical to virtual. Virtuality of noumenon (thing-in-itself, Kant’s Ding an sich) is always flexible enough, compared to phenomenon, to camouflage the on- going adverse environmental processes, the byproducts of the unlimited and unregulated profit- oriented human activities. Nature consumption is accelerated by the guiding ideology of the un- leashed economic man in the sensible world of the living creatures. The sensual reality of accelerated extinction is effectively masked by the virtual sciences of the abstract: ideas, numbers, data, statistics, algorithms, equations and models.

The present taxonomic scenario is disappointing. Over-discussed questions of nature and natural hierarchies remained unanswered or even obscured in a genuine phylogenetic perspective by reams of virtual DNA clades. Answers are misled and manipulated by ideological contexts: what are species, what are subspecies and what are the problematic races? Despite of Darwin’s desperate trials, the ranking and organisational hierarchies remained contradictory. Placing discrete bounda-

T

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ries on the continuous process of diversification in the universe has produced endless debate, especially in the human created realms of subspecies and races. The product oriented nature-exploitive and competitive western culture has significantly influenced the process-oriented and more nature- cooperative eastern cultures and getting world- wide dominance by globalization. Destroying nature resources and ecological services are emerg- ing symptoms of modern western ideology. These simple symptoms are distracted by “green” movements to such euphemistic slogans like “ecological footprint” in order to camouflage the ideological reality of consumption-idiotism behind: why /how we accelerate the rate of nature consumption. During this permanent “progress” most resources have been removed from the taxonomy, from the only integrative science to answer directly and openly these questions. Funds are chan- nelled and disposed either to genetics or to the social projects of evolution. As a result, our taxonomy remained mostly stuck in the century old pathway of macromorphology and intentionally unarmed by the lack of modern revisions, syn- opses and monographs.

Taxonomic state in the Potamophylax cingulatus species group

No progress has yet been realised in the taxonomic application of the empirical resources of the fine phenomics. This huge innovative potential of taxonomy has been left without human and financial resources. Its intrinsic and innate empirical nature is almost suppressed by the piles of virtual neutral molecular markers. But science has self-generating innovative power acting even in such a neglected discipline like taxonomy as has been presented by Szczesny (1990) and Moretti et al. (1994).

Here we sample and apply some theoretical achievements of the phylogenetic species concept to a particular creatures of caddisflies with unsettled taxonomy. One of the initial fine phenomic approaches to caddisfly taxonomy was realised in the Potamophylax cingulatus species group by comprehensive comparative studies on the fine

structures of the phallic organ. Stable shape divergences have been discovered both in the aedeagus and the paramere structures and independent taxa have been discussed, but the possi- bility of species polymorphism was considered at least by a question mark (Szczesny 1990). High polymorphism was suggested again, but not documented in a recent study (Martinez et al. 2016), and the historical polymorphism being a sympatric phenomenon has been debated in the Potamo- phylax genus (Oláh 2017). In a detailed study on the fine structure of the aedeagus and the para- meres the polymorphism was not supported and the shape divergences exhibiting high stability as well as coupled with allopatry permitted to delineate and to describe several new subspecies in the Potamophylax cingulatus group (Moretti et al.

1994).

The discovery of the selective/adaptive speciation trait (Oláh et al. 2015, 2017) has initiated concentrated research first (1) on the fine structure and function of the caddisfly intromittent organ as well as (2) on the structural organisation of periphallic organs, especially the paraproct. A- mong the periphallic organs the paraproct (intermediate appendages) is the structure more inti- mately involved in the cryptic female choice during the copulation processes. These selective traits proved to be sensitive enough to detect early stages of reproductive isolation serving the function of reproductive barriers delimiting incipient species of the unified phylogenetic species concept.

In this paper (1) we review briefly how the unified phylogenetic species concept has been e- volved; (2) how to take subspecies and race out of science; (3) how it is applicable to the taxonomy of the Potamophylax cingulatus species group;

and (4) why the previously supposed polymorphism and the still existing taxonomic rank subspecies (or race) in reality represent independent incipient sibling species. However, based on our theoretical considerations (Oláh et al. 2017) our first motivation was to examine and to convert the subspecies status, still unsettled in the stenophylacini tribe, to phylogenetic sibling species status in this caddisfly group.

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THEORETICAL PART Species concepts

Contemporary systematics is getting to refor- mulate the taxonomic practices by a demanding perspective to delimit and to describe taxa based on phylogenetic history. Yet, non-phylogenetic and non-history based species concepts, like the biological species concept, still remains popular.

Species concepts should not conflict with evolutionary history, but often do. Therefore, it is reasonable to argue for the necessity of a phylogenetic species concept (Velasco 2008). Species is a confusing homonym with three meanings: (1) name of a taxonomic rank (a level or rank in Lin- naean hierarchy, a taxonomic category); (2) word to a particular taxon of that rank, (ontological category, different kinds or ways of being); (3) word to the concept of an evolving group of organisms. This ambiguity is disparate ontologically, but related semantically (Hey et al.

2003). Confusion arises often between the species as taxa, (groups of organisms with shared set of traits) and the species as evolving group of closely and multiple related individuals.

Species concepts serve two disciplines: taxonomy and evolutionary theory (Cracraft 1987).

Accordingly the term species has two basic functions: (1) the species category as a rank in the Linnaean hierarchy created by taxonomist for grouping organisms and (2) the species as taxa with a location in space and time and referring to objective, observable entities, to living objects perceptible by touch (Mayr 1996). Species are dynamic, evolving individuals, almost like a quantum systems but human attempts to force them into rigid classes. Species are real evolutionary groups as well as the human-made categories created by subjectively perceived distinction. The neo-Darwinian synthesis treated the biological species ambiguously as real or subjectively delimited, discrete or nondiscrete, irreducible or decomposable into smaller units depending on particular groups of organisms. How to maintain the unity and discreteness of species in the Darwinian evolutionary transformations

along the branches? How entities can be discrete and still transform over time? These difficulties can be alleviated if species are defined in terms of evolutionary process, as a product of evolutionary phenomena incorporating small genetic changes and the mechanism of natural selection (Cracraft 1987) or rather an alternative idea of integrative organisation (Oláh et al. 2017).

Species is not real. The old linear view of species evolution driven by mutations, recombi- nation and selective pressure and producing a distinct product of species is slowly replaced by a more complex reality of species differentiating, diverging, merging and reverting while driven by diverse integrative mechanism against external and internal impacts. As a result, most of the species categorization applied by taxonomists is inherently and obligatory arbitrary (Hunter 2006).

Many believe that species rank does not exist;

it is not a real category in nature. Darwin doubted the distinction between species and varieties thinking that species is indefinable in spite of the title of his book “Origin of Species”. Despite scepticism over the species category, there are pragmatic reasons for keeping the word species:

the species taxa that are the groups of organisms are real (Ereshefsky 2010). Many genetic studies have re-examined taxonomies of various groups of organisms based on morphology and frequently uncovered paraphyletic or polyphyletic groupings, confirming or refuting previous interpretations.

Studies on mitochondrial DNA diversity conclud- ed that mtDNA data and traditional morphological taxonomic assignments tend to converge (Avise &

Walker 1999). The same data have been revisited with an opposite conclusion (Hendry et al. 2000):

the mtDNA discontinuities do not match recognised taxonomic species. Species realities have been questioned, species category abandoned and new descriptive scheme was suggested for grouping organisms by specifying the amount of differences in various traits at any levels of the phylogenetic tree of life. This conclusion was independent of the marker types used to identify discontinuities. It was interpreted by fundamental flaws in the species paradigm. Today it is clear

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that expectation to find any direct correspondence between neutral markers and adaptive phenomic splits is a naïve unfounded reductionist trial (Oláh et al. 2015).

Only species is real tangible objects. Origi- nally Dobzhansky (1935) has given undisputable ontological significance to the biological category of living individuals. Later (1937), while bringing the Mendelian genetics and the Darwinian evolution together, he has drawn the attention that species are the most stable units in taxonomic practice, as compared either with infraspecific categories such as variety, race, subspecies, or supra-specific ones such as genus, or family.

During this Modern Synthesis species was treated as fundamentally different entity from taxa of higher and lower levels in the hierarchy of biological organisation. According to this misleading concept only the species taxon is the product of evolution, functioning in a direct way as gene pools; exist as whole, as real things (Mayr 1942, 1963). The term species refers to a phenomenon of the nature; species are concrete describable objects. Contrary to species, higher or lower taxa were viewed as subjective and arbitrary, not as an existing real entity (deQueiroz 1985). In the New Systematics the species definable as distinct self- perpetuating units with an objective existence have a greater reality in nature, as dynamic evolving entities that exist independently of human observer. Species have a greater degree of objec- tivity, than higher taxonomic categories which are not definable in this concrete way (Huxley 1940).

This view is still survived repeating that ranking above or below species level is more subjective and ranks above species are relational, lacking the biological reality of the species (Claridge 2010).

Besides questioning the reality of higher taxa along the species tree in the name of modern synthesis, the new systematics has produced more severe disaster with long lasting consequences culminating today in the biodiversity epoch. Mo- dern Synthesis has started to undermine the science of taxonomy by giving priority to experiments, statistics, ecology and genetics and down- graded the empirical descriptive and comparative

nature of taxonomy. Literally suggesting that “an increase in the scientific staffs of the museums is urgently needed if they are to escape from the burden of routine description and naming” of species (Huxley 1940 p. 38). After the new systematics arrived to replace taxonomy most of the available funds moved to genetics in the name of taxonomy. The second disaster came to taxonomy in the present biodiversity epoch when museums are intentionally converted to "baby-sitter centres"

instead of regaining their real function of collect- ing, describing and naming species before their extinction. The core mission of taxonomy is to collect, discover, describe and classify units of biodiversity, the living companies of the human being.

All phylogenetic taxa are real tangible objects.

In his phylogenetic systematics Hennig (1950, 1966) has radically changed this ontological controversy created by the New Systematics of the Modern Synthesis (Dobzhansky 1937, Huxley 1940). He has incorporated the role of evolution in understanding and formulating higher taxa.

Similarly to species the higher level taxa are real, tangible product of evolution. They exist above species level as monophyletic groups composed of the constituting ancestral species, a complete system of common ancestry, an adequate clade, and as the natural outcome of the process of evolutionary descent. The only evolutionary significant property of higher taxa is whether they com- prise this monophyletic clade or not. Genera and families exist as a whole of complete monophyletic clades, outside of the mind of taxonomists (deQueiroz & Donoghue 1988). Higher taxa are real and no any level in the hierarchy is biologically more significant than any other. The weakness of treating species and higher taxa together is that species boundaries are delimited by theoretically well supported qualitative methods, and in contrast, boundaries of higher taxa are subjected to quantitative study, and their patterns is not explained adequately due to lack of theories (Barraclough 2010). Yes, in studies on the evolution of biodiversity the species are the fundamental evolutionary units. From the very beginning of life history studies huge primary

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practical and secondary theoretical data has been accumulated in their phenomics by empirical comparative observations of qualitative nature.

Qualitative phenomics in taxonomy are self-ex- planatory, like in fractal languages or in medical treatment strategies, including cancer failures and cognitive reflections work (Oláh et al. 2015, 2017) Contrary, higher taxa are intensively studied mostly quantitatively by algorithms and models as well as by never tested presumptions, thought experiments.

Velasco (2008) gave crucial demonstrative role to phylogenetic tree to understand phylogenetic inferences. Trees help to visualize important concepts such as what a monophyletic group is and how it is constituted by an ancestral and all of its descendants or how two species are reciprocally monophyletic having all haplotypes of one species more closely related to each other than any haplotypes from his sibling and vice versa.

Tree thinking makes easy to understand how recency of common ancestry, not morphological (morphological species) or interbreeding (biological species) similarity, that defines genealogical relationships. Besides giving real tangible existence to higher taxa on the tree of life, the phylogenetic systematics has initiated a theoretical transformation or rearrangement of the outdated biological species concept into the phylogenetic species concept.

Biological species concept. Darwin (1859) has replaced the Platonic idea and the Aristotelian typological “form” or “essence” concept of species based on type specimen by the evolutionary species concept of the lineage segment, “branches in the lines of descent”. However, the old tradition of species category remained intact functioning further as a rank in the taxonomic hierarchy and predetermined a species concept with fixed temporal and spatial stage, an adult stage at the artificial time-slices of lineages instead of dynamic lineage or branch along the line of descent.

Challenged by the spreading ideas of the phylogenetic systematics the discrete boundaries of the

“adult” biological species on the continuous process of diversification along branches of lineage

segments has produced endless debate and developed multitudes of species concepts (Mayden 1997). The essence of the widely accepted biological species is the discontinuity created and maintained by reproductive isolation representing groups of interbreeding natural populations that are reproductively isolated from other such groups (Mayr 1996). The short definition of the biological species concept is: “Species are groups of interbreeding natural populations that are reproductively isolated from other such groups”.

However, the species criterion of reproductive isolation is not applicable to the reticulate evolution, to organisms with asexual reproduction as well as to the classification of fossil organisms.

The ontology of the biological species concept is incorrect. It lacks generality, not applicable to asexual organisms and inextensible in time. An evolutionary analysis demands temporal extensi- bility. The evolutionary phylogenetic species concept has been formulated and started to challenge the spatial and temporal blindness of the biological species concept. The naïve full-fledged biological species concepts of the New Systematics and the Modern Synthesis have retarded taxonomy upon the morphologically well separated architecture of the “adult” biological species with reproductive isolation. The lack of perfect reproductive isolation is the reason why a subspecies, although distinct morphologically, are not a biological species. Biological species concept cannot be applied to the temporal dimension of species;

unable to specify precisely the limits of species in time; not sensitive enough to recognise adequately the phylogenetic incipient species. Biological species represent a fixed stage of evolutionary divergence; a stage in the evolutionary stream where interbreeding groups of individuals became segre- gated and split into two or more groups incapable to interbreed (Dobzhansky 1937). Large geogra- phically subdivided populations or polytypic biological species often comprising multiple evolutionary entities with or without evolutionary cohe- sive interbreeding. These entities are inherently ambiguous, difficult to demarcate clearly even with intensive field research and applying pro- babilistic threshold with the classic “75% rule”.

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Biological species and its focus on reproductive isolation is a product of the Modern Syn- thesis, but in fact conflicts with much of the cur- rent evolutionary thought and distorts history.

Reproductively isolated groups might be non- monophyletic and creating problems in phylogenetic tree building in diferent ways (Velasco 2008). (1) Paraphyly problem: biospecies can be paraphyletic composed of some, but not all, of the descendants of some ancestral population; there are two populations, one than splits, one of the splitted lineages becomes reproductively isolated from all the others. (2) No tree problem: further speciation events within this paraphyletic biospecies makes real tree building nonsensical. (3) Wrong tree problem: further speciation events within this paraphyletic biospecies produces wrong tree. The history of reproductive isolation, ecological divergence or morphological divergence of speciation events does not define evolutionary history.

The widespread and dominating biological species concept (Mayr 1942) is not in accordance with the new findings that reproductive barriers are semipermeable to gene flow and species can differentiate despite on-going interbreeding (Hausdorf 2011). Biological species concept lumps well differentiated species that nevertheless interbreed regularly. In the unified phylogenetic species concept the species category is being decoupled from the hierarchy of taxonomic ranks and transferred to the hierarchy of biological organisation (deQueiroz 2011). In the old concepts the species as a rank was accepted only if its lineage had reached a particular stage in the process of divergence. Externally allopatric or intrinsically (internally) isolated sympatric (functional allopatry) populations may show every degree of divergence up to that of “full” species (Wilson &

Brown 1953). Lineages that had not yet reached that stage were ranked as subspecies, semi-species or named whatever, like form, variety or race.

Biological species are not comparable entities.

The polytypic species contain a variable number of subspecies, well differentiated evolutionary units or arbitrary subdivisions of continuous spatial variation others include only one monotypic species. For Darwin the distinction along the lineages, lumping or splitting, was unimportant,

because polymorphic variants, clinal variations, forms, geographic races, subspecies, con-species, incipient species and “good” species formed a continuum, the “branches in the lines of descent”

(Mallet 2007).

The phylogenetic species. Species are irredu- cible discrete groups of countable individuals with reproductive cohesion (not disjunction) delineated by heritable diagnostic characters through space and time and exposed to patterns and processes of evolution along the branches in the line of descent. The phylogenetic species is the smallest irreducible, but diagnosable monophyletic group of individual organisms; the smallest set of lineages descended from a common ancestor possess- ing derived, apomorphic traits with unique evolutionary history that is with parental pattern of ancestry and descent (Cracraft 1987). Phylogene- tic species concept is typological in the sense that it is relying upon diagnostic characters in delineation. In the phylogenetic species concept the evolutionary relationships dominates over fertility, contrary to the groups of reproductively isolated interbreeding populations of the biological species concept. If species splitting has not yet reached diagnosability or reproductive cohesion the cluster of species is in statu nascendi (Dozhansky &

Spassky 1959). To rely on reproductive cohesion instead of disjunction is rather reasonable since species and individuals of different higher taxa are frequently interbreeding. Grizzly and polar bear breed in nature (Mallet 2008) and intergene- ric hybrids are well documented among fishes (Burkhead et al. 1991, Garrett 2005), snakes (LeClere et al. 2012) birds (Graves & Zus, 1990, Graves 2007), and primates (Jolly et al. 1997).

Interbreeding of closely related sibling species seems to be a general phenomenon in speciation processes induced along secondary contact zones.

Interbreeding is rather a rule and not a coi- ncidence or exception, under the control of reproductive cohesion and corrected by reinforcement and character displacement.

In our taxonomic modal analysis on caddisflies the entity of phylogenetic species diverged or diverging by fine structures of the reproductive

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barriers, defined by specific initial split criterion, and detected by the degree of morphological dif- ference as an indication of the underlying degree of reproductive isolation. The phylogenetic incipient species is recognised by the diagnostic characters of speciation traits. This is the structure representing reproductive barrier of the biological species concept as well as manifesting potential negative fitness effects in copulating processes. In this way the phylogenetic incipient species concept focuses on the earliest stages of speciation.

Adaptive speciation trait to separate and describe species has been successfully applied recently in detecting, delineating and describing over two hundred caddisfly siblings (Oláh et al. 2012, 2015, 2017, Oláh & Oláh 2017), combining in practice the essence of the phylogenetic and biological species concepts: initial split by reproductive isolation.

How to delineate phylogenetic species? There is inherent subjectivity in all kind of species delineation, like in any kind of entity delineation down to quantum level. In most research fields, but particularly in quantum physics and in human behavioural research the observation has a direct effect on the outcomes (Hey et al. 2003). Under- standing reality is limited by the capacities of observer, by his mental processes and influenced by his interest. Every cogniser has a different relative being of anything. Even the “absolute beings” could be observed from infinity of Nietzsche’s perspectives and could be described by infinity of potential properties or aspects (Oláh et al. 2017). Similarly debated is the role of taxo- nomists playing in the creation of species taxa by taxonomic rank designation.

Species delimitation is frequently confused with species conceptualization. This results in controversy concerning definition of species category and the methods to detect species boundaries. The primary species criterion of the separately evolving metapopulation lineage is widely accepted for species conceptualization. According to this general lineage species concept species are segments of population-level lineages. There is however disagreement about the various secon-

dary species criteria, the operational species criteria, those no longer considered relevant to species conceptualization but only to species delimitation that is to assess lineage separation: (1) intrinsic reproductive isolation, (2) diagnosability, (3) monophyly (Queiroz 2007a, b). Most contemporary species concepts are consistent with the idea that species are evolving lineages or evolving populations. Taxonomic uncertainty is rooted in the evolutionary nature of species; therefore it is unlikely to be solved completely by standardiza- tion (Isaac et al. 2004). Many diverging orga- nisms are still able to mate and produce viable offspring, frequently in contact zones. Changing environment may accelerate divergences on ecological time scales of hundreds or a few thousands of years reinforced by character displacement, reaching a point of no return. Contrary there are convincing cases for reverse speciation where lineages seemed to converge again; with an in- creasing number of hybrids speciation may go into a reverse, reaching a point of separation reunite (Hunter 2006).

An epistemological problem remains however, how to delineate species in space and time along these continuously changing lineages? It might be very difficult to assess empirically a particular taxon. Taxonomist’s tools, circumstances, including sensual and mental capacities and personal interest influence the weight to be given to neutral or adaptive traits and to their particular pattern of variation in designating and describing new species taxa. Taxonomic entities are evolutionary and demographically dynamic, often not very distinct and can change over time or regularly in contact zones (Hey et al. 2003). Moreover, boundaries of all entities are sharp or fuzzy depending upon the spatial and temporal scales of detection that is on the spatiotemporal point of view of the observer (Cracaft 1987). Species, genera and families represent different nested monophyletic clades with temporal scales of separations. They are tangible taxa integrated on population level in the groups of individuals inside of these nested monophyletic clades and along the time course of phylogenetic divergences.

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How to establish fixed stages for any taxa in the dynamic evolutionary stream of processes permanently working over incipient species, ma- ture species, or incipient genera? Taxonomist’s question is what criteria help to identify species taxa? Evolutionist’s question is what criteria aid best to discover locations, boundaries and properties of evolutionary entities? Finding initial split criteria of the phylogenetic species concept may help to answer both questions. Discovering initial split helps to draw the lines of demarcation among evolving entities. The essence of the phylogenetic species delineation is to recognise the first empirical (and/or genomic) sign of the early stages of reproductive barrier building in reproductive cohesion (not disjunction!). Due to ephemeral stages of the continuous process of differentiation and the lack of widely accepted easy or obvious thresholds indicating when speciation has been completed, that is an oversimplified detection of initial splits is troublesome (Winker 2010).

The problem of possible infinite division. Final argument against the phylogenetic species concept refers to the theoretical and practical possi- bility of the infinitely fine divisions for initial splits to differentiate among diverging groups of reproductive cohesion. With whole-genome analyses any two individuals become diagnosably different and could be supported by different monophyly. Character/gene trees and organismal trees are controversial and contradictory: taxa can be monophyletic for one character and non-monophyletic for another and cladograms are really

“cloudograms” superimposed by lineage reticulation. The testable, therefore objective diagnosability and monophyly can be found at any level of hierarchy, but question remains where to draw the lines between lineages? Diagnosability and the smallest cluster depend on the resolution power of the character analyses. Diagnosability and reciprocal monophyly, that is the monophyly with respect to each other, could be produced by extinction of intermediate forms (Zachos &

Lovari 2013). With enough traits all individuals are diagnosable from each other.

The apparently infinite division is further supported as well as distracted by the reductionist

assumption incorporated in all algorithms and models of phylogenetic reconstruction, both by DNA sequences and by unrooted phenetics of numerical taxonomy, that divergence (splitting the lineages) occurs in nature, not reticulation (melding of lineages). But in nature reticulation (the bête noir for cladistics) dominates over divergence and integration over selection according to the general organisational system: aggregates of element in interaction (Botnariuc 1967). Both the reticulation and divergence, like the nature itself, are organised in fractal pattern occurring in the largest and in the smallest, irreducible cladistics units.

Fractal is the nature‘s geometry and organises itself by the negentropy of integration, reticulation against the entropy of disintegration, divergence and selection. Introgression type of reticulation, by melding of lineages, tends to generate phylogenetic discordance more effectively among closely related groups of species, unlike lateral gene transfer. The amount of gene flow by introgression and reticulation of hybridisation is vastly underestimated (Mallet et al. 2015).

Finite division by speciation super traits. The common vernacular argument against phylogenetic species is that every single organism is genetically and phenetically unique. Yes, like every quantum in the Universe! No, because like every quantum, while trying to integrate itself to maintain its integer state, is transformed finally into new emergent entity of natural kind powered by the organising forces of integration (Oláh et al.

2017), and balancing around an idea expressed as nominal kind. Emergence is the appearance of a new observable that cannot be derived from the root theory (Longo et al. 2015). Only reductio- nism, like phenetic species concept in taxonomy and phenetic clade construction in systematics believes that a system can be reduced to the sum of its part. In organizational systemic hierarchy diversification is based on emergence of new entities and the emergent properties differ from those of the constituent subunits (Botnariuc 1967).

Similarly, species as emergent entities are not divisible infinitely into smaller units. Several protective mechanisms evolved in time to produce stable emergencies and to defend their produced

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integrity. Species level organisational emergency cannot be subdivided further if the produced entities of the initial splits are delineated by adaptive traits of the reproductive barrier. In this case the shared derived characters of monophyletic clades are the adaptive structure itself which is creating and maintaining the reproductive isolation. Further subdivision is highly resisted by selection or sexual integration, and the introgression of hybridization may occur without strongly affecting the genomes. But genomic admixture of reticulation nevertheless is realised if the introgressed alleles are established. The adaptive structures of the initial split are stable and highly protected.

The initial split is a symbol for a dynamic temporal dimension representing the genesis of lineage, the splitting of lineage, the birth of a new lineage entity. Initial split is recognised by operational criteria of the various species concepts during the delimitation process of the splitted entities. The splitted is a real entity in nature, a phylogenetic, evolutionary lineage. Species are entities that form lineages or lineage-forming biological entities (deQueiroz 1999). Split entities are gradually becoming more and more differentiated;

reproductively incompatible, ecologically distinct, phenetically distinguishable, diagnosable, and reciprocally monophyletic. Depending on the different contemporary species concept and adopting their different priorities for properties of species delineation, disagreement and conflicts are inevi- table as well as group specific, how to recognise exact temporal splitting of the separately evolving lineage. Species are clusters of organisms passing a threshold of divergence determined by one or several operational criteria. Thresholds for each operational criterion should be fixed by experts of disciplines under the principle of avoiding over- splitting. However, threshold finding should not be realised by numerical or mathematical evaluation systems and neither by putting together unjustified operational criteria, like adaptive shape divergence and neutral DNA markers under the name of multi-source integrative taxonomy (Seifert 2014).

Initial split of diverging species could be recognised not only by detecting direct signs of reproductive isolation or presenting other phylogenetic branching events, but simply empirically by the rarity of hybrids and intermediates between clusters and species (Mallet et al. 2015). These adaptive structures of initial splits are the speciation super traits frequently detectable only by fine phenomics (Oláh et al. 2017). However, in routine observation the speciation super traits seem stable and subtle products of adaptive speciation processes integrated in allopatric isolation and their stability is organised and maintained by several integrative and protective genomic mechanisms (Oláh & Oláh 2017). These protective mechanisms may create nonlinearity in the effect of primary gene flow, or in the secondary one across contact zones, on the processes of divergences, especially in the genomic building of reproductive barriers. This is why even at high rates, gene flow cannot prevent speciation driven and established by adaptive traits of reproductive barriers.

In delimiting the smallest diagnosable cluster of individual organisms there is focus on phenotypic evidences setting aside genetic data (Tobias et al. 2010): (1) proper nucleotide data are not yet sufficiently available; (2) what is available has no relation to the adaptive structures of initial splits; (3) no widespread agreement on how nucleotide data can be used to delimit species. Examining larger portion of the genome to pinpoint specific genes associated with the observed phenotypic differences of the initial split (Patten 2010) seems not very promising to answer the basic questions how to detect initial splits in speciation. There are no well-defined genes, in the sense of the traditional Mendelian term, exist behind the traits of the initial splits (Oláh et al.

2017). There is however, thousands of sequences with almost infinite combinations of pleiotropic, epistasis and epigenetic mechanisms behind mi- nor shape divergences. Frequently they are unde- tectable empirically, diagnosable only with virtual geometric morphometrics. It seems that the adaptive, therefore stable and subtle shape divergen-

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ces, establishing a reproductive barrier, are created and supported by very complex genomic processes. Moreover protein-coding sequence con- vergence in the early branches of the tree of life and high level of incomplete lineage sorting in contemporary divergences make lineage delineations challenging even with whole-genome analyses (Jarvis et al. 2014).

Underestimated biodiversity. Without recog- nizing phylogenetic incipient species the biodiversity is much underestimated by relying upon the outdated and overly lumped alpha taxonomy of

“adult, “full” or “good” species (Pratt 2010). The traditional subspecies concept identifying mini- mum diagnosable units in allopatry as terminal taxa could be essentially synonymous with the phylogenetic species concept (Remsen 2010).

This is clearly confirmed indirectly by the findings that in a meta-analysis of molecular phylogenetic monophyly (Zink 2004) only 3% or in a new global meta-analysis (Phillimore &

Owens 2006) around 36% of avian subspecies represent distinct phylogenetic lineages as measured by the neutral mitochondrial DNA marker. This is not surprising, because neutral markers are unable to measure adaptive traits of subspecies involved in the initial splits while building the reproductive barriers. Much geographic variation may arise via selection;

therefore, DNA tests restricted to selectively neutral genetic data are misleading, neutral markers are not associated directly and firmly with local adaptation. Selection yields distinct phenotypes invisible to neutral markers (Patten 2010). In spite of these finding mixed teams of taxonomists and geneticists remained on the old pathway of trying to couple any kinds of phenotypes with routine neutral markers. They are lucky if, by accident, shapes and neutral sequences fit to each other. If not, Procrustes superimposition starts working. Our distinction between non- adaptive neutral and adaptive non-neutral morphological traits demonstrates that neutral markers are rather blind and not sensitive enough to detect the real on-going adaptive selection processes, that is the adaptive molecular mechanisms creating the divergences on relevant loci pro-

ducing the speciation traits in the early stages of speciation (Oláh et al. 2015, 2017). The lack of congruence between phenotypic traits and neutral molecular data, particularly at sibling species or at subspecies level (Cicero 2010) is very indicative.

It refers to adaptive processes triggering and governing diagnosable traits just at or around the initial splits. The evidence of the detected overall incongruences directly suggests that subspecies could be incipient phylogenetic species, representing the early stages of speciation (Mayr 1942, Phillimore 2010). Moreover, a genuine consensus about subspecies concept is difficult to achieve, because trinominal epithets may cover hetero- geneous mix of evolutionary phenomena and cannot be classified as strict science in the fuzzy world of realism (Fitzpatrick 2010).

Taking subspecies and race out of science Unified species concept. After fundamental theoretical studies deQueiroz (2007a, b) has suggested a unified species concept. He has clearly distinguished and separated the causal processes that produce the lineages (how nature works!) and the operational criteria used to recognize them in practice (how human ranks!). Different species concepts are just tools of the taxonomists in order to find species in their various lifecycles along the stages of speciation. He has retained the general concept of species as separately evolving metapopulation lineages that is the causal process, the only necessary property of species. All the other properties are treated as contingent properties and treated as necessary for considering lineages to be species: phenetically distinguishable, diagnosable, monophyletic, intrinsically reproductively isolated, and ecologically divergent. These properties remain important first (1) as operational criteria to delineate species as evidences of lineage separation for the existence of species and second (2) to define subcategories or recognise different classes of species precisely, based on the given properties: reproductively isolated species, diagnosable species, monophyletic species, ecologically differentiated species. This clear separation of the conceptual problem of defining species category from the methodological problem of

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species delimitation helps research by focusing disagreement to species delimitation with a more demanding perspective for searching species boundaries. The shift in the conceptualization of species category in the unified species concept reducing species criterion to the separately evolving metapopulation lineages has a number of consequences for taxonomy: (1) undifferentiated and undiagnosable lineages are species; (2) all evolutionary lineages, both distinct and indistinct, are species; (3) accepting the integrative frame- work of unified species concept, biologist must regard lineages that merit recognition of species;

(4) morphologically indistinct “cryptic” lineages are diagnosable by other operational criteria (Naomi 2010); (5) species can fuse; (6) species can be nested within other species lineages; (7) species category is the old taxonomic rank; (8) a shift from viewing species category as one member of the hierarchy of taxonomic ranks to viewing it as a natural kind whose members are the units at one of the levels of biological organisation; (9) encouraging taxonomist to develop new methods of species delimitation (deQueiroz 2007a, b); (10) shift from classifying organisms to testing hypotheses about lineage boundaries and phylogenetic relationships (deQueiroz 2005).

However, the reality of this shift, from describing species to phylogenetic studies, is unjustified; al- together over 100 million (Lee 2016) or including prokaryotes one to six billion (Larsen et al. 2017) species is waiting to be discovered, recognised and described before their extinction.This unified species concept was working behind and influenced our studies to discover the speciation super traits as a new method of species delimitation for initial splits, as well as helped us to recognise and to describe over two hundred incipient caddisfly species during a few years, mostly in the sky islands of the so called well studied European mountain ranges (Oláh et al.

2015, 2017, Oláh & Oláh 2017). Moreover, if we go into the details and study its roots and its postmodern background philosophy, the unified species concept applies a refined fuzzy version of the old essentialism, going back to Plato and Aristotle.

Fuzzy essentialism. We have been devising and using taxa from the very beginning, ever

since our ancestors evolved the capacity for language, on an essentialist basis of species. This was enforced later by Platonic and Aristotelian essences and killed recently by Darwin, who has fostered, rather than settled questions about what species really are. There is untapped information in our mind and in our language: species are categories of natural kinds (Hey 2001). However, evolutionary biologists are more interested in the entities of evolutionary groups and not in the mental contributions to taxa. The natural kinds with perceived degree of distinction are based on their essences represented and manifested by varying individual entities. The evolutionary groups might or might not be distinct in space and time, capable of myriad ways of gene exchange to create groups within groups over time. The species problem is fostered by the conflicting motivations to recognise categories of natural kinds with real essences and to understand evolutionary groups. Anti-essentialist critiques are often misplaced and unproductive (Haslam 1998).

We have to remember that entities in the set theory are (1) crisp, deterministic, and precise in characters; (2) dichotomous of yes-or-no, rather than more-or-less; (3) and dual of true-or-false, rather than in between. But complexity of entities increases along organisation of natural kinds. Our ability to make precise statements becomes almost mutually exclusive, both ontologically and epis- temologically. Probability and uncertainty theories have been developed to model these uncer- tainties of reality. Fuzzy set theory is one of these theories, generated to exceed dual logic of clas- sical set theory in order to understand continuity and discontinuity in the ever-changing structural reality starting from quantum sets to sets of living entities.

The world is a collection of objects, assorted into types (Kitcher 2007). In the ontology of biological entities the taxa are natural kinds with real essences of balancing equilibrium underlined by variability ranges of character states including hidden microstructure that scientifically disco- verable, essential to the kind, and making the kind what it is. Ideas, concepts and categories are nominal kinds. The natural kinds are contrasted

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with nominal kinds following Locke’s distinction between the real essences of characters that par- titions the nature into kinds of entities as metaphysical or ontological reality and the nominal essence of abstract ideas, definitions or categories mediated by human concepts. Natural kind is ontic structural realism (1) how entities are organised in nature. Nominal kind is epistemic construal (2) how human ranks entities. The ever changing clinal essentialism of natural kinds, as contrasted with Plato and Kant, is composed of distinct components (Haslam 1998): (1) core of necessary properties; (2) inherent or intrinsic hidden structures underlying superficial properties in supervenience; (3) determinate extensions even with vague boundaries defined by these properties; (4) underlying intrinsic properties are causally related to the accessible characters; (5) despite developmental transformations and graded variation the essential sameness is stable; (6) great inductive potential with wide variety of inferences and generalisations.

Historical concept of race. The concept of race divides Homo sapiens into a small number of groups based on some type of (1) biological foundation, (2) discrete racial grouping, (3) inhe- ritance, (4) genealogy of geographic origin, and (5) physical phenotypes. Conceptual, ontological, epistemological and normative controversies have been accumulated due to ambiguities and con- fusions generated during race boundary delineation; due to moral status of racial identity and solidarity; due to justice and legitimacy of poli- cies; due to institutions and aimed at undermining racial inequality (James 2017). Three competing schools of thought form three metaphysical camps. (1) Racial naturalism holds the old biolo- gical conception of race bearing biobehavioral essences with underlying natural heritable genetic and phenetic properties explaining behavioural, characterological, and cultural predispositions of individual entities of racial groups. (2) Racial constructivism holds that even if biological race is false, races exist through human culture and human decisions. (3) Racial scepticism of elimi- nativists holds that races of any type do not exist and racial naturalism is false and recommends discarding the concept of race entirely.

Metaphysics of race or subspecies. Biological research on race motivated by or lending credence to underlying racist attitude created great pains for scientists to deny the existence of biological human race. Nevertheless, human races adapted to particular environments do in fact exist (Pigliucci

& Kaplan 2003). Already Voltaire wrote, well- packed with Locke’s empiricism, that only blind people could doubt that there are different races (subspecies). People, like any other living creatures, can be classified according to their differences detected, experienced, measured and described in taxonomical studies by various traits of gross morphology, fine phenomics or genetic structure. There are emergent entities exist, like phylogenetic species in spite of speculative trials to formulate arguments against the reality of biological races from blind (neutral) genetics, relativity, and anti-racism. Natural kind is a group of objects characterised by some trait-variability equilibrating around objective essence that is the mind-independent similarity. Social kind is a group of objects with similarity based in existing social practices, institutions, or conventions. So- cial construction is a classification whose members constitute a social kind. In different sense, but biological realists and social constructivists agree about the reality of race. However, they disagree about the kind of racial categories:

biological realists say race is natural kind; social constructivists say race is social kind. Elimina- tivists say: there are no races; racial attributions are false; race is neither biologically real nor socially real. Social constructivists and elimina- tivists agree that races are not natural kinds, but they disagree about the reality of races. Social constructivists admit that race is real even though it is not grounded in genetic differences. Elimina- tivists are error theorist claiming that race is an empty term; nothing belongs to this category;

conditions of race criteria are not satisfied by anything (Diaz-Leon 2012).

Taxonomist or evolutionist, the competent au- thors and users of the species, subspecies, and race concepts maintain that natural property is a necessary condition of taxa. There is no scientific ground for social constructionist view. But this

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view could still be defended by semantic exter- nalism, simply spoken, by support from outside.

In conceptual analysis the semantic externalist in- sights from the critique of the analytic/synthetic distinction can be extended to justify social constructionist position (Haslanger 2006). Moreover, if conditions of natural property satisfied, the naturalism of the biological realists was the correct view. If these conditions not met the view of error theorists were correct.

Philosophical debate on the semantics of general terms and on criteria for real kinds is widen- ing. An obscure concept of basic racial realism to escape the defeat of antirealist position was ela- borated by formal logic applying and combining plethora of notions, all packed into the ontological suitcase: social kind, real kind, real social kind, scientifically relevant kind, unkind, kindred, ro- bust kind, basic kind, genuine kind, basic realism, unkind realism, scientific realism (Glasgow &

Woodward 2015). The concept of basic racial realism was intended (1) to provide an exciting and powerful resource for thinking about race; (2) to capture useful and applicable parts of race that we need to make social progress; (3) do not deny that the features that make races are biological features; (4) to decide races still by visible, biological features, not by social properties; (5) but it does not commit to there being real biological races, that fit poorly with ordinary race-talk; (6) avoiding moral disasters that have plagued racial characterization throughout modern history. Basic racial realism suggests in one metaphysical way, that human beings look just different and sorting us into different categories, but those categories are neither biological kinds nor socially dependent kinds. As a result race is neither biologically real nor socially real, it is real all the same, but most important that the new concept camouflages the anti-science byproduct of this tragicomic debate over reality of race, whether race is biologically real, socially real, or simply not real.

Social construction of human race. Social con- struction of race is realised by impersonal and personal agents highly exposed to contingent choices. Impersonal causal agents (cultures,

conventions, institutions) construct by previous visual-conceptual experiences, by powerful prior notions, by background theories, by nonrepresen- tational phenomena. Personal social agents construct through their choices determined or influenced by scientists’ judgement like theory selection, experiment evaluation, as well as by personal interest/power relations. Shift in human classification has been documented to follow the shift of interest and power (Mallon 2014). What is constructed by these agents, the human traits or human kinds, are designed by culture rather than by biology or nature. These agents construct human traits by evaluating inferences from very complex and contradicting social influences in theory production and from the social construction of facts with ungrounded scientific rationality, scientific realism or scientific process (Laudan 1981, Nelson 1994). In contrast, naturalist attitudes towards science are based (1) on epistemological fundamentalism of empiricism and causal modelling; (2) on metaphysical fundamentalism of supervenience and reduction governed by natural laws; (3) on human naturalism of nonanomalism and methodological naturalism (Mallon 2014).

Races are incipient species! Negating natural kind of human races ignores the basic achievements of modern biology (Mayr 2002). In spite of the social and political connotations there is a naturalistic approach gathering strength to stop the social destruction of race (Sesardic 2010).

Yes, but there is a sound potential for a scientific destruction of the race! The unified species concept gives a real perspective to take race out of phylogenetics, human genetics and taxonomy.

Race is the incipient phylogenetic species, which is the basic concept of Darwinism. There is however, a cost to overcome the century-old debate about the role of race in science. But this cost is not as high as compared to the recently suggested liberal solution to take race simply out of science by slimy substitution dictates in the name of de- mocracy and use of terms like “ancestry” or “population” to describe human groupings. They say that language matters also in racial thinking (Yudell et al. 2016). Instead of this anti-science